Graduate Thesis Or Dissertation
 

Structural and functional characterization of candidate vernalization genes in barley

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  • Vernalization - the requirement of a period of low temperature to induce the transition from a vegetative to a reproductive state - is an evolutionarily and economically important trait in the Triticeae. The genetic basis of vernalization in barley (Hordeum vulgare subsp. vulgare), a model crop for the Triticeae, was first defined in terms of a three-locus epistatic model. Candidate genes for two of the vernalization loci (Vrn1 and Vrn2) were recently cloned in diploid and polyploid Triticurn species. The sequence and expression of HvBM5A, a candidate for Vrn-H1, were characterized in a panel of ten barley germplasm accessions, including one accession of the wild progenitor (Hordeum vulgare subsp. spontaneum). Differences in vernalization requirement are most likely due to allelic variation at the HvBM5A promoter and/or intragenic sites rather than in the coding region. An analysis of promoter and intron 1 polymorphisms showed that differences in the sequence of the latter are most consistent with growth habit classifications. HvBM5A expression patterns correlate with growth habit and meristem development. HvBM5A maps to the predicted position of Vrn-H1 on chromosome 5H. All available evidence suggests that HvBM5A is Vrn-H1. The presence/absence of the tightly linked ZCCT-H gene family members on chromosome 4H is perfectly correlated with growth habit in the germplasm array: all spring forms show a complete deletion of the ZCCT-H gene family. All available evidence suggests that a ZCCT-H gene family member is Vrn-H2. All QTL for vernalization requirement reported for barley in the literature can be explained by a two-locus model involving the Vrn-H1 and Vrn-H2 candidates. The data from this study provide a rigorous definition for "facultative growth habit": facultative genotypes have the Vrn-H2 deletion and signature "winter" habit alleles at Vrn-H1. These data lay the foundation for determining if the coincidence of Vrn-H1 with QTL for multiple winter hardiness-related phenotypes is due to linkage and/or pleiotropy.
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